A STUDY OF THE REMAINS OF CROCUTA FROM
THE KÜÇÜKYOZGAT DISTRICT
MUZAFFER SÜLEYMAN ŞENYÜREK, Ph.D.
Professor of Anthrppology and Chairman of the Division of Palaeoanthropology University of Ankara
During the course of the excavations1 I carried out in the fossiliferous beds of the Küçükyozgat ( E l m a d a ğ )2 district in the summer of 1954, was found a fragment of an upper jaw and a fragment of a lower jaw belong-ing to the genus Crocuta3. These fossil remains were found in a deposit of hard, heterogeneous calcareous clay of light brown color at the Kuyu larla locality (Locus 2), 4 which I had discovered during the course of field work and excavations in the Küçükyozgat region in 1-951.5
In my previous studies,6 I had designated the fauna of whitish cal careous marls of lacustrine origin, occurring to the south of Sarıkaya Ağılı (the corral of Sarıkaya), "Küçükyozgat fauna" to distinguish it from the fauna of the heterogeneous calcareous elays of light brovvn color 7 from Kuyu tarla and Örücebağ,8 which I had called "Karacahasan - fauna".
1 I wish to extend my thanks at this time to the Office of the Dean, the Professors' Council and the Central Anatolian Research Station of the Faculty of Language, History and Geography of the University of Ankara for the grant to carry out excavations in this region in the summer of 1954. Gn this occasion I also wish to thank Prof. Hamit Nafiz Pamir, Geaeral Director of Maden Tetkik ve Arama Enstitüsü (Institute for
Mineralogical Research) of Ankara, for providing me a jeep in the last phase of my field work in the "Küçükyozgat district.
2 For the location of this fossiliferous region see Tschachtli, 1942, p. 323; Şenyürek, 1953 a, p. 1; Erol, 1954, fig. 1.
3 During the course of the 1954 exeavations in this region fragmentary remains of a.hyaenid were also found in whitish calcareous marls of lacustrine origin at Küçük yozgat, at the new fosilliferous locus (for this see Şenyürek, 1953a, fig. 3), I had located in 1951, which is just to the southeast of the locus first visited by Tschachtli (Tschachtli, 1942, p. 325) in 1941 (for Tschachtli's locus see Şenyürek, 1953a, fig. 2). I have deferred the study of these remains from this locus until further material from this whitish calcareous marl series is obtained.
4 I have called the Kuyutarla site shown in fig. 1, Locus 1, and the plaee where the remains of Crocuta have been found (figs. 2-3), occuring just to the west of Locus 1 and toward Örücebağ, Locus 2. There are also fragmentary remains of a couple of carnivores from Locus 1, which are stili indetermined (see. Şenyürek, 1953a, p. 2).
5 For the history of researches made at this fossiliferous region, first visited by Tschachtli in 1941, see Şenyürek, 1953 a, pp. 1-2, and Şenyürek, 1953 d, pp. 459-460. 6 Şenyürek, 1953a, p. 2; Şenyürek, 1953b, p. 142; Şenyürek, 1953 d, pp. 459-460. 7 Erolcalls these of pinkish color. See Erol, 1954, p. 93
As the remains of Crocuta found at Kuyutarla have a bearing on the age of the Karacahasan beds, which occur to the southwest9 of the whitish calcareous marls of Küçükyozgat, and as these remains show some interesting morphological variations, I have decided to describe them separately and in detail.
FAMILY HYAENIDAE GRAY, 1869 1 0
SUBFAMILY HYAENINAE MIVART, 1 8 8 21 1 GENUS CROCUTA KAUP, 1 8 2 8 1 2 CROCUTA EXIMIA (ROTH AND W A G N E R ) 1 3
The species Crocuta eximia (Roth and Wagner),1 4 which had originally been described as Hyaena eximia by Roth and Wagner1 5 is represented by a fragment, including the left premaxilla and a large part of the left maxilla (figs. 4-6) and a fragment of right corpus mandibulae (figs. 7-11), found in a deposit of hard, heterogeneous calcareous clay at Locus 2 at Kuyutarla.
In the fragment of the upper jaw are preserved the three incisors, the canine and the four premolars. The palate behind a point corresponding to the middle of the third lobe of the fourth upper premolar is broken and the first upper molar is not preserved. In the right corpus mandibulae the incisors, the canine, the root of P1, the crowns of second, third and fourth premolars and most of tbe crown of the first molar are preserved. The second and third incisors are retained intact. On the other hand, only a part of the crown of the first incisor (figs. 12-13) is preserved, and, as its basal part is missing, it could not be fitted to the preserved root of this tooth. This corpus mandibulae was not found in association with the fragment of the upper jaw, although very close to it. In b o t h t h e upper and lower jaw the teeth are only very slightly worn.
Among the Pontian localities of Anatolia, the existence of the species Crocuta eximia (Roth and Wagner) has so far been reported from only Muğla and Eşme in western Anatolia 1 6 and Kayseri in central
Ana-9 For the location of fossiliferous loci see Erol, 1954, fig. 2. 1 0 Simpson, 1950, p. 117.
1 1 Ibid., p. 118. 1 2 Ibid., p. 118.
1 3 Pilgrim, 1931, p. 116. 1 4 Ibid.
1 5 Wagner, 1860, p. 120; Gaudry, 1862, p p . 80-81; Pilgrim, 1931, p. 116.
1 6 See Ozansoy (1951, p. 148), a n d Yalçınlar (1946, p. 126). In addition, in his study of t h e Pontian beds of the upper Gediz region, in western Anatolia, Yalçınlar (1947, P. 174) states: "Parmi les fossiles Carnivores, il y a une mandibule gauche avec trois
MUZAFFER ŞENYÜREK 30
CROCUTA FROM THE KÜÇÜKYOZGAT DISTRICT
tolia.1 7 In the preliminary reports on these other Pontian localities, only the name of this species has been recorded, without any description of the morphological features of the specimens, and no photographs have been published. For this reason it has not been possible for me to com-pare the remains from Kuyutarla with these other Anatolian specimens.
The Upper Faw : In the fragment of the upper jaw from Kuyutarla the incisors increase in height and size from the first toward the third incisor (see Table ı). The buccal surfaces of all three incisors, in mesial and distal views, are seen to curve conspicuously in the lingual direction. The first and second upper incisors are nearly bicuspid with two tuber-cles at the bottom of the lingual surface, of which the medial one is larger. In the first incisor the distal basal tubercle is less conspicuous than that of the second upper incisor. In the form of the basal tubercles these two teeth of the Kuyutarla specimen do not differ much from those of Crocuta eximia (Roth and Wagner) from Pikermi1 8 and those of Crocuta variabilis (Zdansky) from the Pontian of China.1 9 The first and second upper inci sors exhibit a small marginal tubercle on the distal surface of the crown, which is larger in the second than in the first incisor.
The third upper incisor is a large and caniniform tooth that projects far below the level of the second incisor, as is characteristic of the genus Crocuta.20 The morphology of the third upper incisor of the specimen from Kuyutarla resembles that of Crocuta eximia (Roth and Wagner) from Pikermi2 1 and also that of Crocuta variabilis (Zdansky) from China.2 2 Zdan sky describes the upper third incisor of Crocuta variabilis ((Zdansky) as fol-lows: I3 ist sehr Krâftig, Eckzahn-artig, mit je einer Kante an der Vorder-innen-und Hinter-aussenseite, die durch einen Basalwulst verbVorder-innen-unden sind." 23 This des cription also fits the I3 of the Kuyutarla specimen (fig. 5) and that of Cro cuta eximia (Roth and Wagner) from Pikermi.2 4
A comparison of the meâsurements of the specimen from Kuyutarla
premolaires et une canine rappelant un Hyemde et un autre fragment de mandibule gauche port-ant une premolaire." However, as no photographs are given, it is not possible to assess the identity of the fossil form represented.
17 See İzbırak and Yalçınlar, 1951, p. 156 (according to their statement the speci-fic determinations were made by Yalçınlar). Regarding the attribution of these re mains to Crocuta eximia, İzbırak and Yalçınlar (1951, p. 156), however, state that this
is a "determination provisoire."
18 See Gaudry, 1862, pl. XIV, fig. 1. 19 See Zdansky, 1924, pl. XX, fig. 2.
20 See Gaudry, 1862, pl. XIII, fig. 1; Zdansky, 1924, p. 95 and pl. XX. fig. 1 and pl. XXI, fig. 3; Pilgrim, 1931, p. 114 and pl. I, figs. 1 and 3.
21 See Gaudry, 1862, pl. XIV, fig. 1.
22 See Zdansky, 1924, pl. XX, fig. 2 and pl. XXI, fig. 4. 23 Zdansky, 1924, p. 95.
24 See Gaudry, ,1862, pl. XIV, fig. 1.
32 MUZAFFER ŞENYÜREK
(see Table ı) with the figures given by Gaudry, shows that the upper inci sors of the Kuyutarla specimen are only slightly smaller than those :of Crocuta enmia ( R o t h and Wagner) from Pikermi.2 5 JRobustness values of the upper incisors of Crocuta variabilis (Zdansky),Crocuta honanensis (Zdan-sky) and Crocuta gigantea (Schlosser) from the Pontian of China, calculated from the figures given by Zdansky2 6 and Schlosser2 7, are as follows :
Number of
Specimens Average Range Crocuta variabilis (Zdansky) I1 62 8 4 2 . 8 5 3 3 . 1 2 - 4 8 . 1 4
[from Zdansky, I2 62 9 65.90 61.38- 77.00
1924> P- 97] I3 83 0 149.21 131.25-174.00
Crocuta honanensis (Zdansky) I1 13 1 39.42 —
[from Zdansky, I2 13 2 61.10 —
1924 and Schlosser, 1903] I3 23 3 119.90 90.00-149.80
Crocuta gigantea (Schlosser)
[from Schlosser, 1903] I3 1 143.00 —
From these figures it is seen that the upper incisors of the Kuyutarla specimen (see Table 1) are, in size, in the range of Crocuta variabilis (Zdan sky) and are larger than those of Crocuta honanensis (Zdansky). The third upper incisor of the Kuyutarla specimen also exceeds that of Crocuta gigantea (Schlosser) from China whose other teeth are extremely large (see the tables).
The upper canine of the Kuyutarla specimen is separated from the third upper incisor by a diastema whichis about 7.0 m m . wideat thebase. In norma lateralis the tip of the third upper incisor comes almost to the level of the tip of the canine. However, when the canine is examined care-fully, it is seen that the enamel margin of the buccal surface is stili con-siderably above the alveolar margin (fig. 4), indicating that the canine had not yet completed its eruption, although all the other upper perma-nent teeth had fully erupted. It is thus evident that when the canine had
25 Gaudry (1862, p. 86) gives the following figures as "diametre principal" of the the upper incisors of Crocuta eximia (Roth and Wagner) from Pikermi: I1 9.0.mm.,12
10.0 mm., I3 15.00 mm. 26 Zdansky, 1924, p. 97. 27 Schlosser, 1903, p. 34 and p. 36. 28 Specimens 1, 8, 10, 19, 20 and 24. 29 Specimens 1, 8, g, ıg, 20 and 24. 30 Specimens 1, 2, 3, 8, g, 18, 19 and 24. 31 One specimen given by Zdansky, 1924. 32 One specimen given by Zdansky, 1924.
CROCUTA FROM THE KÜÇÜKYOZGAT DISTRICT 33
fully erupted, its tip would project considerably beyond that of I3, as is the case in Crocuta eximia (Roth and and Wagner) from Pikermi 34 and Crocuta variabilis (Zdansky) from China. 35
The upper canine from Kuyutarla presents a well developed anterior and an equally well developed posterior edge. Zdansky has described the upper canine of Crocuta variabilis as follows: "Der Canin hat zwei Lângskanten, welche die flache Innenseite von der stark konvexen Aussenflache trennen." 36 This description also fits the specimen from Kuyutarla and that also of Crocuta eximia (Roth and Wagner) from Pikermi. 3 7
As the basal part of the crown is stili embedded in the alveolus, its length and breadth measurements could not be taken. The buccal height, however, could be measured, owing to a fracture in the middle part of the external wall of the alveolus (see Table 1). Arambourg and Piveteau have published the picture, stated to be natural size, of the upper jaw of a specimen of Crocuta eximia (Roth and Wagner) from the Pontian of Salonika, which is in a younger stage of development than our specimen.3 8 As far as can be judged from the photograph, the height of this upper canine from Salonika appears to be shorter than that of the Kuyutarla speci men. In Specimen 8 of Crocuta variabilis (Zdansky) from China, depicted by Zdansky, 39 the upper canine is well preserved. In this specimen of Crocuta variabilis (Zdansky), as far as can be judged from the photograph, the height of the upper canine comes close to that of the Kuyutarla specimen.
The upper first premolar of the Kuyutarla specimen is separated from the upper canine by a diastema which is about 6.0 mm. wide at the base. As it is, the specimen from Kuyutarla differs considerably from the speci-mens of Crocuta eximia (Roth and Wagner) from Pikermi 40 and Samos H and from Crocuta variabilis (Zdansky) from China, 42 in all of vvhich the space separating P1 from C1 is much shorter. However, as was stated before, the canine of the upper jaw from Kuyutarla had not yet com-pleted its eruption. Thus when the canine had fully erupted this diastema would probably become much shorter and approach in width those of Crocuta eximia (Roth and Wagner) and Crocuta variabilis (Zdansky), which resemble each other in having a short diastema between P1 and C 1.
34 See Wagner, 1860, pl. 3, fig. 9 and Gaudry, 1862, pl. XIII, fig. 1. 35 Zdansky, 1924, pl. XX, fig. 1 and pl. XXI, fig. 3.
36 Ibid., p. 95.
37 See Gaudry, 1862, pl. XIV, fig. 1.
38 See Arambourg and Piveteau, 1929, pl. X, fig. 2. in this picture the buccal surface of this upper canine from Salonika is clearly visible
39 See Zdansky, 1924, pl. XX, fig. 1.
40 See Wagner, 1860, pl. 3, fig. 9 and Gaudry, 1862, pl. XIII, fig. 1 and pl. XIV, fig- ı.
41 Pilgrim, 1931, pl. 1, fig. 1.
42 Zdansky, 1924, p. 95, pl. XX, figs. 1-2 and pl. XXI, figs. 1-4.
3 4 M U Z A F F E R Ş E N Y Ü R E K
As far as can be judged from the drawing published by Gaudry, the first upper premolar of the specimen from Kuyu tarla seems to be less dis-placed lingualwards than in the specimen of Crocuta eximia (Roth and Wagner) from Pikermi. 43 In this feature, Crocuta variabilis (Zdansky) from China comes closer to the Anatolian specimen. 44
The first upper premolar of the Kuyutarla specimen has a single cusp and a single root and in its general morphology it does not differ nluch from that of Crocuta eximia (Roth and Wagner) from Pikermi4 5 and Cro cuta variabilis (Zdansky) from China.4 6 Zdansky describes the first upper premolar of Crocuta variabilis (Zdansky) as followsc "Seine Krone ist im Umriss fast Kreisrund. Von der stumpfen Spitze zieht eine Kante nach hinten und nach vorne, welche die konvexe Aussenflache von der in vertikaler Richtung konkaven Innenflache trennt." 47 This description would almost fit the specimens of Crocuta eximia (Roth and Wagner) from Kuyutarlâ (fig. 5), and also, as far as can be judged from the picture published by Gaudry,4 8 that from Pikermi.
From Table 3 it is seen that in size, as expressed by the robustness value, the first upper premolar of Kuyutarlâ specimen falls within the range of variation of Crocuta eximia (Roth and Wagner) from Europe, but exceeds the maximum of Crocuta variabilis (Zdansky), which on the aver-age has a smaller first upper premolar than Crocuta eximia (Roth and Wagner). On the other hand, P1 of the Kuyutarlâ specimen is smaller than that of Crocuta honanensis (Zdansky) and is far exceeded by that of Crocuta gigantea (Schlosser). It is of interest to note that the first upper premolar of Lycyaena dubia Zdansky is smaller than the minima of all the Pontian species of Crocuta listed.
The crown index of the first upper premolar from Kuyutarlâ is lower than those of all the available specimens of Crocuta eximia (Roth and Wag-ner) and Crocuta variabilis (Zdansky). The crown index of the first upper premolar from Kuyutarlâ is very near that of a specimen from China, originally described as Hyaena sp. by Schlosser,49 which was subsequently attributed to Crocuta honanensis by Zdansky,50 but is lower than that of the type specimen of the latter species and than that of Crocuta gigantea (Sch losser). Lycyaena dubia Zdansky is characterized by a relatively low crown index, which is considerably below that of the Kuyutarlâ specimen.
4 3 Gaudry, 1862, pl. -XIV, fig. 1.
4 4 Zdansky, 1924, pl. X X , fig. 2, and pl. X X I , figs. 2 a n d 4. 4 5 Gaudry, 1862, pl. X I V , fig. 1.
4 6 Zdansky, 1924, pl. X X , fig. 2 a n d pl. X X I , figs. 2 and 4. 4 7 Ibid,. p. 95.
4 8 Gaudry, 1862, pl. X I V , fig. 1. 4 9 Schlosser, 1903, p. 34.
CROCUTA FROM THE KÜÇÜKYOZGAT DISTRICT 35
The second upper premolar of the Kuyutarla specimen, separated from P1 by a very short diastema, possesses a main middle cusp a small posterior tubercle, followed by a talon behind, and a relatively speaking well developed anterior tubercle which is located on the mesio-lingual corner of the crown (figs. 5-6).
In size the second upper premolar from Kuyutarla falls within the range of variation of Crocuta eximia (Roth and Wagner) from Europe and Crocuta variabilis (Zdansky) of China (Table 4). P2 from Kuyutarla is, like the other Pontian species of Crocuta, greatly exceeded in size by that of Cro cuta gigantea (Schlosser), but surpasses that of Crocuta honanensis (Zdansky) which, however, falls within the range of Crocuta eximia (Roth and Wag-ner) and Crocuta variabilis (Zdansky). In size the second upper premolars of Lycyaena chaeretis (Gaudry) and Lycyaena dubia (Zdansky) fall far below the minimum of the Pontian species of Crocuta listed. In crown index P2 from Kuyutarla is within the range of variation of Crocuta eximia (Roth and Wagner) and Crocuta variabilis (Zdansky). P2 from Kuyutarla exhib-its a higher crown index than that of Crocuta honanensis (Zdansky) and Crocuta gigantea (Schlosser). It is seen that Lycyaena chaeretis (Gaudry) and Lycyaena dubia Zdansky are distinguished from all the Pontian specimens of Crocuta listed in having a relatively narrower second upper premolar. The third upper premolar of the Kuyutarla specimen is in contact with the second upper premolar and possesses a main cusp, a well devel oped anterior tubercle and a posterior tubercle (fıg. 5) .)This tooth dif-fers from the second upper premolar mainly in being larger and in pos-sessing a larger and higher posterior tubercle and in having a moderately developed anterior cingulum which is in front of the anterior tubercle.
In possessing a relatively well developed anterior tubercle the second and third upper premolars of the Kuyutarla specimen resemble the corres-ponding teeth of Crocuta eximia (Roth and Wagner) from Pikermi.5 1 Pil-grim states that in the second and third upper premolars of Crocuta eximia
(Roth and Wagner) from Pikermi an anterior tubercle "is invariably pre-sent". 52 Kittl has published the picture of a third upper premolar of Cro cuta eximia (Roth and Wagner) from Maragha, which also shows a well developed anterior tubercle. 5 3 On the other hand, in the second and third upper premolars of Crocuta variabilis (Zdansky) from China, as stated by Zdansky, an anterior tubercle is completely lacking in some specimens while it is present in others. 5 4 Thus the second and third upper
premo-51 See Gaudry, 1862, pl. XIV, fig. 1.
52 Pilgrim, 1931, p. 117. From the photograph of the upper jaw of Crocuta eximia (Roth and Wagner) from Salonika, published by Arambourg and Piveteau (1929, pl. X, fig. 2), it can be seen also that the third upper premolar possesses a large anterior tubetcle.
53 See Kittl, 1887, pl. XVII, fig. 2.
3 6 M U Z A F F E R Ş E N Y Ü R E K
lars of Kuyutarla specimen approach those forms of Crocuta variabilis (Zdan-sky) exhibiting an anterior tubercle, and differ from specimens of this form from China which are devoid of such a tubercle. Zdansky states that the second upper premolar of Crocuta honanensis (Zdansky) possesses an anterior tubercle. 55 In the photograph of a third upper premolar from the Pontian of China, published by Schlosser and described by him as Hyaena sp., which may belong to Crocuta honanensis (Zdansky) is also seen a well
developed anterior tubercle. 56 Thus in this feature the Kuyutarla speci men also does not seem to differ from Crocuta honanensis (Zdansky). A small anterior tubercle is observed also in the second and third upper pre-molars of Crocuta gigantea (Schlosser). 57
As can be seen from Table 5, in size the specimen from Kuyutarla is within the range of variation of Crocuta eximia (Roth and Wagner) and Crocuta variabilis (Zdansky). This tooth of the Kuyutarla specimen is slight-ly larger than the corresponding tooth of Crocuta honanensis (Zdansky), which is withir the range of Crocuta eximia (Roth and Wagner) and Crocuta variabilis (Zdansky) but is greatly surpassed by that of Crocuta gigantea (Schlosser). The robustness values of the third upper premolars of Lycyaena chaeretis (Gaudry) and Lycyaena dubia Zdansky again fall far short of the minima of the Pontian species of Crocuta. In crown index the third upper premolar from Kuyutarla falls again in the range of variation of Crocuta eximia
(Roth and Wagner) and Crocuta variabilis (Zdansky). The crown index of the Kuyutarla specimen is lower than those of Crocuta honanensis (Zdansky) and Crocuta gigantea (Schlosser) which exceed the maximum value of Cro cuta eximia (Roth and Wagner) and fall within the upper part of the range of Crocuta variabilis (Zdansky) in this index. The crown index of the third upper premolar of Lycyaena chaeretis (Gaudry) is lower than the minimum of Crocuta variabilis (Zdansky) and is near that of Crocuta eximia (Roth and Wagner), being only very slightly higher than the minimum of this spe cies. On the other hand, the crown index of the third upper premolar of Lycyaena dubia Zdansky falls short of the minima of Crocuta eximia (Roth and Wagner) and Crocuta variabilis (Zdansky).
The fourth upper premolar of the Kuyutarla specimen shows an an terior, a middle and a posterior lobe, which, as can be seen from figs. 4-5, increase in length from before backwards. The third lobe is relatively long, amounting to 40.81% of the total length of the crown. 58 In this
5 5 Zdansky, 1924, p. 104.
5 6 See Schlosser, 1903, pl. I I I , fig. 5. 5 7 Ibid., p. 36 and pl. I I , figs. 3-4.
5 8 As calculated from the figures given by Gaudry, the third lobe of P4 represents 42.50% of the crown length in Crocuta eximia (Roth and Wagner) from Pikermi a n d 38.23% in P4 of Lycyaena chaeretis (Gaudry) again from Pikermi (For the measurements of these forms see Gaudry, 1862, p. 86 a n d p. 94). As calculated from the figures given by Pei (1934 ,p. 107) the third lobe of P4 amounts to 35.14% of the crown length in specimen
C R O C U T A F R O M T H E K Ü Ç Ü K Y O Z G A T D I S T R I C T 3 7
feature this fourth upper premolar differs from that of genus Hyaena, in which the third lobe is relatively short, 59 and agrees with genus
Crocuta, in which this lobe is lengthened. 60 In this tooth from Kuyutarla the protocone, or the mesio-lingual cusp, is strongly reduced (figs. 5-6).
The fourth upper premolar from Kuyutarla, in having a strongly reduced protocone, differs conspicuously from the fourth upper premolars of genus Lycyaena, 61 Crocuta salonicae (Andrews), 62 Crocuta honanensis (Zdan-sky), 63 later, post-Pontian forms of genus Crocuta, 64 and members of ge nus Hyaena, 65 in all of which the protocone is well developed. Crocuta gigan-tea (Schlosser) from the Pontian of China has a relatively small protocone in its fourth upper premolar, 6 6 but stili a comparison with the photograph published by Schlosser 67 shows that the protocone of P4 of the Anatolian specimen is less distinct than that of Crocuta gigantea (Schlosser).
In having a strongly reduced protocone, P4 of the Anatolian specimen resembles that of some forms of Crocuta eximia (Roth and Wagner) and Crocuta variabilis (Zdansky). Regarding the development of protocone in the fourth upper premolar of Crocuta eximia (Roth and Wagner) from Pikermi, Gaudry makes the following statement: "....le talon interne de la carnassiere superieure de l'Hyaena eximia, au lieu d'etre bien developpe comme dans les autres hyenes, est aussi mdimentaire que dans le genre chat. Wagner a dejâ constate cette particularite ; elle est tres frappante sur le crâne que j'ai recueilli." 68 A comparison with the drawing published by Gaudry, 6 9 shows that the protocone of the fourth upper premolar of the Anatolian specimen is as much reduced as in this Pikermi specimen. 7 0 Pilgrim, on the other hand,
C C
a n d to 36.58% in specimen — o f Crocuta crocuta sinensis (described as Hyaena sinensis C.794 C.750
Zdansky by Pei) from the lower Pleistocene of China.
5 9 Gaudry, 1862, p. 88 and Arambourg a n d Piveteau, 1929, p. 124. 6 0 Gaudry, 1862, p. 82 and Arambourg and Piveteau. 1929, p. 124.
6 1 Pilgrim, 1931, p. 103. See also Gaudry, 1862, p. 93 a n d Del C a m p a n a , 1914, pl. X I [1], fig. 2.
6 2 Pilgrim, 1931, p. 115 a n d p. 123. 6 3 Zdansky, 1924, p. 104.
6 4 Gaudry, 1862, p. 88; Pilgrim, 1931, p p . 115-116; Pei, 1934, pl. X I V , fig. ıc a n d pl. X V I I , fig. ı b ; Hooijer, 1952, fig. 3.
6 5 G a u d r y , 1862, p. 88.
6 6 Schlosser, 1903, p. 36 and Pilgrim, 1931, p. 115. 6 7 Schlosser, 1903, pl. I I , fig. 2a.
6 8 Gaudry, 1862, p. 82. 6 9 Ibid., pl. X I V , fig. 1.
7 0 Regarding the protocone of P4 of Crocuta eximia (Roth a n d Wagner) from M o n t Leberon G a u d r y (1873, p. 17) states: "Elle est de meme depourvue de denticule interne." For the protocone of P4 in Crocuta eximia from the Pontian of Salonika, Arambourg a n d Pive teau (1929, p. 123) make the following statement: "Le denticule interne (deuterocone ), place a la partie anterieure de la dent, est mdimentaire; la carnassiere presente par suile un aspect tres coupant."
3 8 M U Z A F F E R Ş E N Y Ü R E K
makes the following statement regarding the protocone of P4 in Crocuta eximia (Roth and Wagner) from Pikermi: "The material of the Pikermi species which I have examined shows a distinct variation in the size of the protocone." 71 Regarding the size of protocone in the fourth upper premolar of the spe cies Crocuta eximia (Roth and Wagner) Arambourg and Piveteau also state: ". . . .mais il est facile d'observer, sur l'ensemble des pieces de Pikermi, de Maragha, etc, de grandes variations dans la taille de ce tubercule." 72 Zdansky states with regard to the protocone of Crocuta variabilis (Zdansky): "Auch P4 variiert, vor Allem durch die Ausbildung seines Innenhöckers recht betrâchüich. Dieser ist stets schwach entwickelt, am schwachsten bei Ex. ı (Taf. XXI, Figg. 3&4 ) und Ex. 20, wo er nicht stârker als bei H. eximia von Pikemi ist, am stârksten bei den Ex. 8 (Taf. XX, Figg. 1&.2 ) und 21." 73 In the size of the protocone, P4 of the Kuyutarla specimen resembles that of Specimen 1 of Crocuta variabilis 74 in which the protocone is strongly reduced, and differs notice-ably from that of Specimen 8 of this form from China 75 in which this cusp is considerably stronger than in our specimen. In short, it is seen that P4 of the Kuyutarla specimen agrees only with the specimens of Cro cuta eximia (Roth and Wagner) and Crocuta variabilis (Zdansky) with strongly reduced protocone, that is with the advanced forms of these.7 6
In size the fourth upper premolar from Kuyutarla falls in the range of Crocuta eximia (Roth and Wagner) ar d Crocuta variabilis (see Table 6). This tooth of the Anatolian specimen is slightly larger than that of Crocuta honanensis (Zdansky), but is greatly exceeded by that of Crocuta gigantea (Schlosser) in size. P4 of Lycyaena chaeretis (Gaudry) and Lycyaena dubia Zdansky is exceeded in size by that of the Pontian species of Crocuta, ex-cepting a specimen from Taraklia.7 7 In crown index the Kuyutarla spe cimen falls within the range of variation of Crocuta eximia (Roth and Wag-ner) and Crocuta variabilis (Zdansky), but falls short of Crocuta honanensis
(Zdansky) and Crocuta gigantea (Schlosser), which in this index, exceed the maxima of Crocuta eximia and Crocuta variabilis. Lycyaena chaeretis (Gaudry) and Lycyaena dubia Zdansky exceed the averages of Crocuta eximia and Crocuta variabilis,78 but fail short of Crocuta honanenesis (Zdansky) and Crocuta gigantea (Schlosser).
7 1 Pilgrim, 1931, p. 117.
7 2 A r a m b o u r g a n d Piveteau, 1929, p. 124. 7 3 Zdansky, 1924, p. 96.
7 4 Ibid., pl. X X I , fig. 4. 7 5 Ibid., pl. X X , fig. 2.
7 6 According to Pilgrim (1931, p. 115 a n d pp. 117-118) the protocone is also relative-ly small in the fourth upper premolar of Crocuta carnifex and Crocuta mordax, respectiverelative-ly from the Chinji a n d Dhok P a t h a n stages of the Siwalik Hills.
7 7 T h e measurements of this Taraklia specimen have been taken from Zdansky (1924, p. 97). As the original paper of K h o m e n k o (1914) is inaccessible to me, it "will not be possible to say anything definite, as it stands, on the very narrow b r e a d t h measurement. 7 8 Lycyaena chaeretis (Gaudry) very slightly exceeds the maxima of Crocuta eximia and Crocuta variabilis.
C R O C U T A F R O M T H E K Ü Ç Ü K Y O Z G A T D I S T R I C T 3 9
The Lower Jaw: In the mandibular fragment from Kuyutarla there are
two large foramina mentalia, arranged one behind the other, of which the
anterior one, located below the mesial root of P
2, is larger than the
pos-terior one, which is situated under the interalveolar septum between P
2and P
3(see figs. 7-8). In having two foramina mentalia the Anatolian
specimen resembles the majority of Crocuta eximia (Roth and Wagner)
from Baltavar, Pikermi and Maragha, which usually have two foramina.
79T h e locations and the relative sizes of the foramina mentalia of the
Kuyu-tarla mandible are the same as those of the mandibles of Crocuta eximia
(Roth and Wagner) from Maragha, depicted by Kittl
80and De
Mecque-nem.
8 1Regarding the foramen mentale of Crocuta variabilis, Zdansky
states: "Betrâchtliche Unterschiede sind mit Bezug auf die For, mentalia zu
Kons-tatieren. Zuweilen ist nur ein solckes vorhanden und liegt dann hinter der vorderen
oder unter der hinteren Wurzel des P
2. In derselben Lage können sich zwei
get-rennte öder oberflâchlich vereinigte Foramina bejinden. Hâufig tritt auch,
manch-mal nur einseitig, eine Gruppe von drei Foramina mentalia auf."
82As Zdansky,
has published the photographs of only two adult mandibles of Crocuta
variabilis,
83which, as far as can be seen, show only one large foramen
mentale, a direct comparison of the location and relative sizes of the
fora-mina between my material and those specimens of Crocuta variabilis,
described as having two foramina, has not been possible.
84T h e Anatolian specimen, in having two foramina mentalia, differs
from Lycyaenachaeretis (Gaudry) from Pikermi, which usually has one
fora-men .
85However, in the specimens of Lycyaena chaeretis . (Gaudry), from
the Lower Plioeene of the Vienna Basin, described by Zapfe, there are
more than one orifice, two of which are under the premolars.
86Regard-7 9 G a u d r y , 1862, p. 8 4 ; Kittl, 1887, p. 3 3 3 ; De M e c q u e n e m , 1925, p. 23. R e g a r d i n g the variation in the n u m b e r of foramina mentalia in Crocuta eximia ( R o t h a n d W a g n e r ) G a u d r y (1862, p. 84) states: "Wagner a dit que toutes les hyenes ont un seul trou mentonnier,
tandis que l'Hyaena eximia de Grece en a deux. M. Suess, dans l'Hyaena eximia de Baltavar, a aussi remaraue deux trous. f'ai fait une semblable observation sur deux mâchoires que j'ai recueillies a Pikermi; mais sur une troisieme, on ne trouve qu'un trou mentonnier."
8 0 Kittl, 1887, pl. X V I I , fig. 1. 8 1 De M e c q u e n e m , 1925, pl. I X , fig. 7. 8 2 Zdansky, 1924, p. 98.
8 3 Ibid., pl. X V I I I , fig. 3 a n d pl. X X , fig. 3. Zdansky (1924, pl. X X I I , fig. 4) has also published the lateral view of a juvenile m a n d i b l e , described by h i m as Hyaena?
vari-abilis, in which t h e nurhber of foramina is not clearly seen.
84 Crocuta honanensis (Zdansky) possesses one large orifice (see Zdansky, 1924, p l . X X I I I , fig. 4 ) , which is located u n d e r the mesial root of P2. However, r e g a r d i n g t h e m e n t a l e foramen of this Chinese fossil, Zdansky (1924, p. 104) states: "Auf der rechten Seite
befindet sich ein grosses For. mentale unter der Mitte des P2. Ob weiter hinten ein zweites vorhanden
war, lâsst eine Beschâdigung nicht entscheiden. Rechts ist eine Gruppe von drei Foramina vorhanden."
8 5 G a u d r y , 1862 p. 9 3 ; . Pilgrim, 1931, p. 110.
8 6 Zapfe, 1948, p. 256 a n d figs. 3a a n d b. R e g a r d i n g a third orifice in m a n d i b l e [A] Zapfe (1948, p. 256) states: "Unter I3 befindet sich ein Foramen mentale."
4 0 M U Z A F F E R Ş E N Y Ü R E K
ing these in mandible [A] Zapfe states: "Unter der Mitte von P2 befindet sich ein grosses Foramen mentale, ein Kleines dahinter unter der Vorderıvurzel des P3. " 87 For mandible [B] he states: "Unter der Hinterwurzel des P2 befindet sich ein grosses, unterhalb P3 ein kleines Foramen mentole."88 It is seen that in the location of foramina mentalia the Anatolian specimen differs from Lycyaena chaeretis (Gaudry) from the Vienna Basin.
In the Anatolian specimen the lower border of corpus mandibulae is nearly straight under P4 and P3 and begins to curve slightly downward under the distal root of P2. The height measurements of this corpus mandibulae are as follows :
Height under P2= 4 2 . 5 0 mm. Height under P3= 40.00 mm.
According to Gaudry8 9 in Crocuta eximia from Pikermi the height of corpus mandibulae under P2 is 33.00 mm., which is smaller than that of the Anatolian specimen. As far as can be judged from the drawing pub-lished by Gaudry9 0 the height under P3 also is smaller in the Pikermi specimen than in the Anatolian mandible. On the other hand, as far as can be judged from the photographs published, provided the scales are correct, in corpus height the Anatolian mandible comes closer to speci-mens of Crocuta eximia (Roth and Wagner) from Salonika 91 and Samos.9 2 Thus it appears that the corpus height of the mandibks of Crocuta eximia (Roth and Wagner) is quite variable and that the greater height seen in the Anatolian specimen does not bar it from inclusion. in this species.93
As far as can be judged from the photographs published by Zdansky, 94 the- corpus mandibulae of the Anatolian specimen is considerably deeper than that of Crocuta variabilis (Zdansky). It appears, thus, that in corpus height the Anatolian mandible comes nearer to those of some forms of Crocuta eximia than to that of Crocuta variabilis.
The lower incisors of the Kuyutarla specimen increase in size from the first toward the third incisor (see Table 2). The lower incisors possess a marginal tubercle, on the distal surface, which also increases from
8 7 Zapfe, 1948, p. 256. 8 8 Ibid., p. 260. 8 9 Gaudry, 1862, p. 86. 9 0 Ibid., pl. X I I I , fig. 2.
9 1 Arambourg and Piveteau, 1929, pl. X, fig. 3. 92 Pilgrim, 1931, pl. I, fig. 1.
9 3 T h e same result also comes from the comparison of the corpus height under M1.
In" the specimen of Crocuta eximia ( R o t h a n d Wagner), studied by G a u d r y , this height is 45.00 m m . (Gaudry, 1862, p. 86). On the other h a n d , in three adult specimens of this species from Pikermi (Nos. 49673, M 8970 a n d M 8971), measured by Pilgrim, this dimension is 50.00 m m . , while in one from Samos (No. M4164), studied by the same author, it is 42.00 m m . (see Pilgrim, 1931, p p . 119-123).
CROCUTA F R O M THE KÜÇÜKYOZGAT DISTRICT 41
the first toward the third incisor.In the first and second lower incisors
of this specimen the bucco-lingual diameter exceeds their mesio-distal
diameter, while the reverse is the case in the third lower incisor, owing
to the relatively larger size of the distal marginal tubercle of this tooth.
In Kuyutarla mandible, in norma lateralis, the incisors are
implanted in front of the lower canine to the same extent as that seen in
Crocuta eximia (Roth and Wagner) from Salonika and Samos, depicted
respectively by Arambourg and Piveteau
95and Pilgrim
96and those of
the mandibles of Crocuta variabilis, published by Zdansky.
97Regarding
the location of I
2in Crocuta variabilis, Zdansky states: " I
2steht etwas hinter
I
1und I
3."
9 8T h e same feature is also seen in the Kuyutarla specimen.
As far as can be judged from the photograph published by Arambourg and
Piveteau,
99in Crocuta eximia from Salonika also the distal surface of I
3seems to be slightly more forward than that of I
2.
The measuremehts of the lower incisors of Crocuta from Kuyutarla
are listed in Table 2. As'for the comparable material, our information on
the dimensions of the lower incisors of Pontian species of Crocuta is limited
to the lower incisors of Crocuta variablis (Zdansky), and to I
3of Crocuta
honanensis (Zdansky) and Crocuta gigantea (Schlosser). Robustness values
of these calculated from the figures given by Zdansky
100and Schlosser
101are as follows :
Crocuta variabilis (Zdansky) I
1[from Zdansky, 1
21924, p. 99] I
3Crocuta honanensis (Zdansky)
[from Schlosser, I
3I903, P. 34]
Crocuta gigantea (Schlosser)
[from Schlosser, 1903, I
3P. 36]
N u m b e r of
Specimens
9.1028
1036
104 1 1Average
20.48
42.25
69.76
7 2 . 0 095.00
Range
19.20-23.56
37.23-50.02
62.56-77.18
—
—
95 Arambourg and Piveteau, 1929, pl. X, fîg. 3. 96 Pilgrim, 1931,-pl. I, fig. 1.
97 Zdansky, 1924, pl. XVIII, fig. 3, pl. XX, fig. 3 and p. 98. 98 Ibid, p. 98.
99 Arambourg and Piveteau, 1929, pl. X, fig. 3a. 100 Zdansky, 1924, p. 99.
101 Schlosser, 1903, p. 34 and p. 36.
102 Specimens 1, 6, 8, 9, 10, 11, 12, 13 and 15. 103 Specimens 1, 6, 8, 9, 11, 12, 13 and 15. 104 Specimens 1, 6, 8, II, 12 and 15.
42 MUZAFFER ŞENYÜREK
It is seen from these figures that in size, I1 and I3 of the Kuyutarla specimen exceed the maxima of Crocuta variabilis, while its I2 falls within the range of this Chinese form. In size I3 of the Anatolian Crocuta exceeds that of Crocuta honanensis, but is greatly surpassed by that of Crocuta gigan-tea (Schlosser).
The lower canine of the Kuyutarla specimen is a well preserved and large tooth. Zdansky describes the lower canine of Crocuta variabilis as follows: "Zwei Langskanten begrenzen an der Innenseite ein schmales, fast ebenes Feld." 1 0 5 This description fits exactly the lower canine of the Kuyutarla specimen (fıgs. 9-11). In the Kuyutarla lower canine the two vertical ridges, or keels, on the lingual surface are well formed.
From Table 8 it is seen that in the length of the lower canine the Kuyutarla specimen falls in the range of Crocuta eximia (Roth and Wagner) from Europe, and surpasses slightly the maximum of Crocuta variabilis
(Zdansky) from China. The Anatolian specimen, in this measurement, also surpasses one specimen of Crocuta honanensis (Zdansky) and the maximum of Lycyaena chaeretis (Gaudry) which, on the average, tends to have a smaller lower canine than the Pontian forms of Crocuta.
In the Kuyutarla mandible only the alveolus of the first lower pre-molar is preserved, showing that this tooth had one root. As can be seen from fig. 10, in the Kuyutarla specimen the alveolus of P1 is separated from the lower canine by a very short diastema, the anterior border of the alveolus of P1 being about 3.5 mm. away from the exposed distal surface of the iroot of the canine at this level. In this feature the Anatolian man dible differs from the specimens of Crocuta eximia (Roth and Wagner)from Pikermi 1 0 6 and that from Maragha, depicted by Kittl 1 0 7 and De Mec-quenem, 1 0 8 in which this diastema is wider. On the other hand, in having a short diastema the Anatolian mandible comes nearer to that of Crocuta eximia (Roth and Wagner) from Salonika, which is described by Aram bourg and Piveteau as follows: "Le diasteme separant la premıere premolaire de la canine est a peu pres nul." 1 0 9 As far as can be judged from the photo-graph published by Arambourg and Piveteau, 1 1 0 the diastema of the Sa lonika specimen seems to be even more reduced than in the Kuyutarla mandible. Thus, the short diastema of the Anatolian specimen is stili with-in the range of variation of Crocuta eximia (Roth and Wagner), which is variable in this feature. In having a short diastema the Anatolian man dible differs from specimens of Crocuta variabilis, stili retaining a P1, in
1 0 5 Zdansky, 1924, p. 98.
1 0 6 Gaudry, 1862, p. 82 and pl. XIII, fig. 2. 1 0 7 Kittl, 1887, pl. XVII, fig. 1.
1 0 8 De Mecquenem, 1925, pl. IX, fig. 7.
1 0 9 Arambourg and Piveteau, 1929, p. 123. See also De Mecquenem, 1925, p. 24. 110 Arambourg and Piveteau, 1929, pl. X, figs. 3 and 3a.
C R O C U T A F R O M T H E K Ü Ç Ü K Y O Z G A T D I S T R I C T 4 3
which, according to Zdansky, the width of this diastema ranges from 5.0 to 8.0 mm.1 1 1 As far as can be judged from the photograph of the man-dible in norma lateralis, published by Zdansky,1 1 2 this diastema seems to be also relatively wide in Crocuta honanensis (Zdansky).
The second lower premolar of the Kuyutarla specimen (figs. 7-9) possesses a main middle cusp, a very small anterior tubercle. and a slightly larger posterior basal tubercle, behind which there is a talon. In having a small anterior tubercle, P2 of Crocuta from Kuyutarla differs from the second lower premolars of the specimens of Crocuta eximid (Roth and Wagner) from Pikermi,1 1 3 Salonika1 1 4 and Maragha, depicted by De Mecquenem,1 1 5 which are devoid of an anterior tubercle.1 1 6 In this fea-ture P2 of the Anatolian specimen resembles that of Crocuta eximia (Roth and Wagner) from Maragha, depicted by Kittl,1 1 7 which shows a tiny anterior tubercle. It also resembles that of Crocuta variabilis, which, accord ing to Zdansky, although variable, often exhibits a small anterior tuber cle.1 1 8 In having an anterior tubercle P2 of the Anatolian specimen differs from that of Crocuta honanensis (Zdansky) in which, according to Zdansky, "ein Vorderhöcker fehlt", 119 which is also true for Crocuta gigantea (Schlosser).120
As can be seen from Table 10, in size, P2 from Kuyutarla falls within the range of variation of Crocuta exîmia (Roth and Wagner) and Crocuta variabilis (Zdansky). It is larger than P2 of Crocuta honanensis (Zdansky), but falls far short of that of Crocuta gigantea (Schlosser). In size of P2, the maximum of Lycyaena chaeretis (Gaudry) falls far short of the minimum of the Pontian species of Crocuta. In crown index P2 of the Kuyutarla speci men falls slightly below the minimum of the available specimens of Crocuta eximia (Roth and Wagner) and comes very near to the minimum of Crocuta variabilis (Zdansky). It is exceeded in this index by Crocuta honanensis (Zdan sky) and Crocuta gigantea (Schlosser), but far surpasşes the maximum of Lycyaena chaeretis (Gaudry). Lycyaena chaeretis (Gaudry), which possesses a 1 1 1 Zdansky, 1924, p. 98. For the specimens of Crocuta variabilis lacking a P1, Zdansky (1924, p. 98) states: "Dieser Zahn fehlt bei Ex. 1 und 12 und gleichzeitig ist das Diastema so kurz, dass er wohl kaum ursprünglich vorhanden war und dann verloren ging, sondern angenommen werden muss, dass er sehon in der Entwicklung ausgeblieben ist."
1 1 2 Zdansky, 1924, pl. X X I I I , fig. 4. 1 1 3 Gaudry, 1862, p. 83 and pl. X I I I , fig. 2.
1 1 4 Arambourg and Piveteau, 1929, p. 123 and pl. X, fig. 3. 1 1 5 De Mecquenem, 1925, pl. I X , fig. 7.
1 1 6 Regarding the anterior tubercle in the lower premolars of Crocuta eximia (Roth and Wagner) Pilgrim (1931, p. 116) states: " P2 and P3 practically without anterior cusps."
1 1 7 Kittl, 1887, pl. X V I I , fig. 1.
1 1 8 Zdansky, 1924, p. 98 and pl. X X , fig. 3. 1 1 9 Ibid., p. 105.
4 4 M U Z A F F E R Ş E N Y Ü R E K
relatively narrow P2, in crown index falls far below the minimum of the Pontian species of Crocuta.
The third lower premolar of the Kuyutarla specimen possesses a main middle cusp, a small anterior tubercle and a larger posterior tubercle with a talon behind it (figs. 7-9). This tooth differs from the second lower pre molar mainly in being larger and in having a more rectangular contour and a somewhat larger posterior tubercle. In this tooth, as well as in P2, the anterior cingulum, on the mesio-buccal corner of the crown, has been reduced. In having a small anterior tubercle, this tooth differs from that of Crocuta eximia (Roth and Wagner) from Pikermi and Maragha, depicted respectively by Gaudry1 2 1 and De Mecquenem.1 2 2 In this feature P3 of the Anatolian mandible resembles that of Crocuta eximia (Roth and Wagner) from Maragha, depicted by Kittl1 2 3 ,and also Crocuta variabilis (Zdansky), some specimens of which show a small anterior tubercle.1 2 4 P3 of the Anatolian specimen differs from that of Crocuta honanensis (Zdan sky), in which the anterior tubercle is negligible1 2 5 and that of Crocuta gigantea (Schlosser), which is, according to Schlosser, devoid of this tubercle.1 2 6
It is seen from Table 11 that in robustness value P3 of the Kuyutarla specimen is within the range of variation of Crocuta eximia (Roth and Wag-ner) and Crocuta variabilis (Zdansky). It exceeds P3 of Crocuta honanensis
(Zdansky) in size, but falls far short of that of Crocuta gigantea (Schlosser). Again, in size of P3, the maximum of Lycyaena chaeretis (Gaudry) is exceeded by the minimum of the Pontian species of Crocuta. In crown index P3 of the Kuyutarla specimen falls within the range of Crocuta escimia (Roth and Wagner), Crocuta variabilis (Zdansky) and Crocuta honanensis (Zdansky). In this index P3 of the Anatolian specimen is also very near that of Crocuta gigantea (Schlosser). In crown index, P3 of Lycyaena chaeretis (Gaudry) falls far short of the minimum of the Pontian Crocuta.
In the fourth lower premolar of the Kuyutarla specimen both the an terior and posterior tubercles are strongly developed. In this tooth the anterior tubercle exceeds the posterior tubercle in size and also in ele-vation. Directly behind the posterior tubercle there is a tiny accessory
1 2 1 Gaudry, 1862, pl. X I I I , fig. 2. 1 2 2 De Mecquenem, 1925, pl. I X , fig. 7.
1 2 3 Kittl, 1887, pl. X V I I , fig. 1. Regarding P3 of Crocuta eximia from Salonika Aram-bourg nad Piveteau (1929, p. 123) state: "P3 et P4 possedent deux tubercules bien developpes."
1 2 4 Zdansky, 1924, p. 98 and pl. X X , figs. 3-4. Zdansky (1924, p. 98), regarding this, states: "Auch hier wiederholt sich der Fall, dass der Vorderhöcker nur durch eine basale Ansch-mellung der Vorderkante des Haupthöckers ersetzt sein kann, z-B. bei Ex. 1 und 12. Bei Ex. 6 ist er unter dem vorhandenen Materiale am Kraftigsten entmickelt."
1 2 5 Zdansky, 1924, pl. X X I I I , figs. 3-4. Zdansky (1924, p. 105) states regarding the anterior tubercle of P3 in Crocuta honanensis : "Ein Basalhöcker an der Basis der Vorderkante is kaum angedeutet."
C R O C U T A F R O M T H E K Ü Ç Ü K Y O Z G A T D I S T R I C T 4 5
tubercle (figs. 7 and 11) on top of the talon, which forms a rather sharp edge on the disto-lingual comer of the crown (figs. 9-11). The anterior cingulum, on the mesio-buccal corner of the crown, is reduced.
In having an anterior tubercle larger than a posterior tubercle, P4 of the Anatolian specimen differs from that of Crocuta eximia (Roth and Wag-ner) from Pikermi and Maragha, depicted respectively by G a u d r y1 2 7 and De Mecquenem,1 2 8 in which, as can be seen from the pictures published by these authors, the anterior tubercle is smaller and also lower in height than the posterior tubercle. On the other hand, P4 of the Anatolian speci men resembles closely a specimen of Crocuta eximia (Roth and Wagner) from Maragha, depicted by K i t t l ,1 2 9 in which also the anterior tubercle is both larger and higher than the posterior tubercle. In this feature P4 of the Kuyutarla specimen resembles also that of Crocuta variabilis (Zdansky)1 3 0 and, as far as can be judged from the photograph published by Zdansky,1 3 1 Crocuta honanensis (Zdansky).
In having a small accessory tubercle behind the posterior tubercle, P4 of the Anatolian mandible, comes closer, among the Pontian forms of Crocuta, to a specimen of Crocuta eximia (Roth and Wagner) from Maragha, depicted by Kittl,1 3 2 in which also an accessory tubercle is seen behind the main posterior tubercle.
Regarding P4 of a specimen of Crocuta eximia from Salonika, De Mec-quenem states: "Le talon de la derniere premolaire presente un denticule interne qui reppelle la 4e premolaire de l'lctitherium hipparionum; ce denticule mangue sur les echantillons de Maragha." 1 3 3 This tubercle is also seen in the speci men of Crocuta eximia from Salonika, depicted by Arambourg and Pive teau 134 and in the specimen of this species from Maragha, depicted by K i t t l .1 3 5 In P4 of the Kuyutarla mandible there is no accessory tubercle
1 2 7 Gaudry, 1862, pl. X I I I , fig. 2. 1 2 8 De M e c q u e n e m , 1925, pl. I X , fig. 7.
1 2 9 Kitti, 1887, pl. X V I I , fig. 1. In the specimen of Crocuta exiınia from Salonika, depicted by Arambourg a n d Piveteau (1929, pl. X, fig. 3 a n d 3a) the anterior tubercle is worn but stili larger t h a n the posterior tubercle. It is possifble that in its fresh condition
the anterior tubercle might also have been higher t h a n the posterior tubercle, as in the Anatolian specimen.
1 3 0 Zdansky, 1924, p. 100, pl. X V I I I , fig. 3 a n d pl. X X , fig. 3. Regarding t h e relative size of the anterior a n d posterior tubercles in P4 of Crocuta variabilis, Zdansky(1924, p. 100) states: " In derselben Linie stehen dann auch Vorder-und Hinterhöcker, jener Krâftiger als
dieser."
1 3 1 Zdansky, 1924, pl. X X I I I , fig. 4. In P4 of Crocuta gigantea (Schlosser), depicted by Schlosser (1903, pl. I I , figs. 7 a n d 7a) the anterior tubercle is worn, b u t it is possible that in its fresh state it was higher t h a n the posterior tubercle.
1 3 2 Kittl, 1887, pl. X V I I , fig. 1. 1 3 3 De Mecquenem, 1925, p. 24.
1 3 4 Arambourg a n d Piveteau, 1929, pl. X, figs. 3 and 3a. 1 3 5 Kittl, 1887, pl. XVII, fig. 1.
4 6 M U Z A F F E R Ş E N Y Ü R E K
on the inner side of the talon, which, on this side, forms a rather sharp and angular edge, as in the specimen of Crocuta eximia from Maragha, de-picted by De M e c q u e n e m1 3 6 and as in some specimens of Crocuta variabilis
(Zdansky).1 3 7
The robustness value of P4 of the Kuyutarla mandible exceeds the maximum of the hitherto available specimens of Crocuta eximia (Roth and Wagner) and falls in the range of Crocuta variabilis (Zdansky), being near the upper limit of the range of this form (see Table 12). In size, P4 of the Kuyutarla specimen stili falls far short of that of Crocuta gigantea (Schlosser). The robustness values of P4 of the Pontian species of Lycyaena fall below the minimum of Pontian Crocuta. In crown index P4 of Crocuta from Kuyutarla is in the range of variation of Crocuta eximia and Crocuta variabilis and somewhat exceeds that of Crocuta gigantea (Schlosser). The crown indices of the Pontian species of Lycyaena are, on the average, lower than those of the Pontian Crocuta. The maximum crown index of Pontian Lycyaena somewhat exceeds the minimum of Crocuta, but falls far short of the maximum of Crocuta from this period.
In the first lower molar of the Kuyutarla mandible, the talonid is broken and missing, the tooth being broken behind the second lobe where it joins the talonid. In this tooth, the first lobe(L=12.7 mm.) is longer than the second (or posterior) lobe ( L = 1 0 . 5 mm.). In the lower part of the distal margin of the second lobe there is no indication whatsoever of a metaconid. 1 3 8 A section of the lower part of the buccal surface of the first lobe is damaged, so it is not possible to determine with certainty whether there was a cingulum in this part or not.
In lacking a metaconid the first lower molar of Crocuta from Kuyu tarla resembles those ot Crocuta eximia (Roth and Wagner) from Pikermi1 3 9 and Salon ika 1 4 0 and Crocuta variabilis (Zdansky) 1 4 1 and Crocuta gigantea
(Schlosser) 1 4 2 from China. In this feature the Anatolian specimen differs from the specimens of Crocuta eximia from Maragha, described by De Mec-quenem, in which there is a metaconid 143 and also from Crocuta honanensis
1 3 6 De M e c q u e n e m , 1925, pl. I X , fig. 7.
1 3 7 Zdansky, 1924, p. 100 a n d pl. X X , fig. 4. Regarding this feature of P4 of Crocuta variabilis, Zdansky (1924, p. 100) states: "Aııf den Hinterhöcker folgt ein Basalwulst, der an seinem (Innenrande meistens eine ziemlich scharfe Kante bildet, die nur bei-Ex. 1 ganz fehlt."
1 3 8 In the first lower molar of the hyaenid from the whitish calcareous marls of Küçükyozgat a small metaconid is observed at this place.
1 3 9 Gaudry, 1862, p. 83.
1 4 0 Arambourg a n d Piveteau, 1929, p. 123. However, regarding the occurrence of a metaconid in the first lower molar of a specimen from Salonika, De M e c q u e n e m (1925, p. 23) states: " . . .nous l'avons distingue sur une carnassiire inferieure provenant de Salonigue
(Coll. Puyhaubert ) . "
1 4 1 Zdansky, 1924, p. 100. 1 4 2 Schlosser, 1903, p. 36. 1 4 3 De Mecquenem, 1925, p. 23.
CROCUTA FROM THE KÜÇÜKYOZGAT DISTRICT 47
(Zdansky), which, according to Zdansky, also posscsses this cusp.
144However, it should be noted here that in the specimen of Crocuta eximia
(Roth and Wagner) from Maragha, depicted by Kittl,
145there is no
indication of a metaconid, showing that Crocuta eximia from this locality
was quite variable in this feature.
T h e length of the first and second lobes of M1 of Crocuta from
Kuyu-tarla is 23.20 mm. T h e total crown lengths and talonid lengths of the first
loyyer molars of Crocuta eximia (Roth and Wagner) and Lycyaena chaeretis
(Gaudry) from Pikermi, given by G a u d r y ,
1 4 6are listed below.
Crocuta eximia Lycyaena chaeretis
(Roth and Wagner) (Gaudry)
Crown length 28.00 24.00
Talonid length 5.00 6.00
Length of first and
second lobes
1 4 723.00 18.00
From these figures it is seen that the length of the first and second
lobes of the first lower molar of Crocuta from Kuyutarla is very near that
of Crocuta eximia (Roth and Wagner) from Pikermi and exceeds that of
Lycyaena chaeretis (Gaudry).
D I S C U S S I O N
T h e foregoing account shows that in most of its diagnostic features
Crocuta from Kuyutarla falls within the range of variation of Crocuta eximia
(Roth and Wagner). Thus the remains from Kuyutarla, consisting of a
fragmentary upper j a w and a fragment of a lower jaw, belong to Crocuta
eximia (Roth and Wagner) which is a characteristic Pontian species.
148It is observed also that in a lârge n u m b e r of features Crocuta eximia
from Kuyutarla resembles Crocuta variabilis (Zdansky) from the Pontian
of China, at least, some specirnens of this form, further bridging the gap
between this Chinese form and Crocuta eximia (Roth and Wagner).
Regarding the affinities of Crocuta variabilis, Zdansky states: "Bedeutende
Âhn-lichkeit besteht dagegen mit H. eximia, deren Verbreitungsgebiet sich pon
Südfank-reich über Süddeutschlahd, Ungarn, Griechenland, Südrussland bis Persien erstreckte.
144 Zdansky, 1924, p. 106. According to Pilgrim (1931, p. 118) this cusp also occurs in the first lower molar of Crocuta mordax Pilgrim from the Siwalik Hills.
145 Kittl, 1887, pl. XVII, fig. 1. 146 Gaudry, 1862, p. 87 and p. 94.
147 Obtained by subtracting the talonid length from the crown length.
148 Arambourg and Piveteau, 1929, p. 124; Pilgrim, 1931, p. 115;Boule and Piveteau, 1935, p. 786.
48 MUZAFFER ŞENYÜREK
Unzweifelhaft steht H. eximia der chinesischen Form sehr nahe. Leider ist jedoch trotz der ziemlich grossen Zahl der bekannten Exemplare nichts veröffentlicht wor-den, mas über die Variationsbreite der Art befriedigenden Aufschluss gibt. Aus den vorliegenden Arbeiten gewinnt man aber den Eindruck, dass Z.B. der Innenhöcker des P4 niemals so gross wird wie an einigen Exemplaren von H. variabilis. Die spâr-lichen Massangâben über H. eximia, die ich in den vergleichenden Tabellen gesam-melt habe, zeigen, dass die Dimensibnen der beiden Formen fast die gleichen sind. Höchstens könnte M1 bei H. eximia etwas lânger sein. Das von Suess (1861 ) aus Baltavâr beschriebene Fossil besitzt einen ungewöhnlich grossen M1, weiters zeigt M1 seine grösste Breite am Vorderlobus und ein sehr schmales Talonid." 1 4 9 It must be rioted in this connection that when Zdansky published his excellent monograph, 1 5 0 there were relatively only a few publications on the mor-phology of Crocuta eximia (Roth and Wagner) and that the important studies of De Mecquenem,1 5 1 Arambourg and Piveteau, 1 5 2 and Pilgrim 1 5 3 had not yet been published. We can now take up the discussion of the points raised by Zdansky.
It is now known that, as has already been stated by Arambourg and Piveteau 1 5 4 and Pilgrim 1 5 5, the size of the protocone of the fourth upper premolar of Crocuta eximia is quite variable, which thus further bridges the gap between Crocuta eximia and some specimens of Crocuta vari abilis. As for the metric features discussed by Zdansky, the measurements of the upper and lower teeth of Crocuta eximia and Crocuta variabilis are contrasted with those of the Pontian species of Crocuta and Lycyaena in Tables 3-13. From these tables it is seen that in size the ranges of Crocuta eximia and Crocuta variabilis greatly overlap and that, on the average, in some teeth Crocuta eximia and in some Crocuta variabilis exceeds the other, while in a few cases the averages of the two forms are very near each other. In regard to the difference in size of M1 mentioned by Zdansky, according to the figures given in Table 13, the average robustness value of Crocuta variabilis is greater than that of Crocuta eximia. A careful scrutiny, however, of this table shows that the relatively small average of Crocuta eximia is caused by an M1 from Samos (No. M4166), given by Pilgrim, 1 5 6 with abnormally small dimensions. As the picture of this isolated tooth is not published by Pilgrim, it is not possible to say whether there has been a recording error or whether this tooth really belongs to Crocuta eximia (Roth and Wagner). Discounting this tooth from Samos, with abnormally small
1 4 9 Zdansky, 1924, p. 101. 1 5 0 Ibid.
151 De Mecquenem, 1925.
1 5 2 Arambourg and Piveteau, 1929. 153 Pilgrim, 1931.
1 5 4 Arambourg and Piveteau, 1929, p. 124. 155 Pilgrim, 1931, p. 117.
CROCUTA FROM THE KÜÇÜKYOZGAT DISTRICT 49 measurements, the average measurements and indices of the first lower
molars of Crocuta eximia listed in Table 13 are as follows157 :
Length = ( 8 ) 27.76 mm. Breadth = ( 5 ) 12.88 mm. Robustness value = ( 4 ) 349.77 mm.
Crown index = ( 4 ) 4 5 - 51 m m .
It is seen, then, that these fıgures are very near the average measure ments of Crocuta variabilis.
It is true that the size of M1 of the specimen of Crocuta eximia from
Baltavâr far exceeds the maximum of Crocuta variabilis in robustness value (Table 7). However, a scrutiny of the figures listed in Table 7 shows that
this exceptionally large robustness value is a peculiarity of M1 of the
Baltavâr form that is not common to the forms of Crocuta eximia from Pikermi and Taraklia. Indeed, the difference in robustness value between
the M 1 of the Baltavâr form and the maximum of Crocuta variabilis is less
than the distance separating the Baltavâr specimen from the maximum of Crocuta eximia from Pikermi. Thus the Baltavâr fossil in having a large
M1, which must be regarded as an individual or local variation, is farther
removed from Crocuta eximia from other European localities than from some specimens of Crocuta variabilis. The average measurements and indices of the first upper molars of Crocuta eximia from Pikermi and Taraklia are listed below :
Length = (4) 6.12 mm. Breadth = ( 4 ) 13-50 mm. Robustness value = ( 4 ) 82.50 mm. Crown index = ( 4 ) 222.56 mm.
It is seen that these average figures come near to those of Crocuta variabilis.
It may be pointed out regarding the talonid of the first lower molar that the size of the talonid in the specimens of Crocuta variabilis, depicted
by Zdansky, 1 5 8 does not differ much from the specimens of Crocuta eximia
from Pikermi, Salonika and Maragha, depicted by various authors. 1 5 9
To the points raised by Zdansky, it may also be added that in exhib-iting a small anterior tubercle in the second and third lower premolars, some (not all) specimens of Crocuta variabilis differ from the European forms of Crocuta eximia. But in this feature, the corresponding teeth of the
mandible of Crocuta eximia from Maragha, depicted by Kittl 1 6 0 and
157 The figures in brackets show the number of specimens.
158 Zdansky, 1924, pl. XVIII, figs. 3-4 and pl. XX, figs. 3-4.
159 Gaudry, 1862, pl. XIII, fig. 2; Arambourg and Piveteau, 1929, pl. X, figs. 3-3a;
Kittl, 1887, pl. XVII, fig. 1; De Mecquenem, 1925, pl. IX, fig.. 7.
160 Kittl, 1887, pl. XVII, fig. 1.
5 0 M U Z A F F E R Ş E N Y Ü R E K
those of the Kuyutarla mandible resemble specimens of Crocuta variabilis exhibiting an anterior tubercle in P2-P3. Crocuta variabilis differs also from the Pikermi form of Crocuta eximia in having an anterior tubercle larger and higher than the posterior tubercle in its fourth lower premolar. In this feature also the fourth lower premolar of the specimen of Crocuta eximia from Maragha, studied by K i t t l1 6 1 and that of the Kuyutarla specimen resemble closely that of Crocuta variabilis. 1 6 2 It is observed that in the features under consideration the same variations occur in both Crocuta eximia and Crocuta variabilis.
The account given above clearly shows that Crocuta variabilis from the Pontian of China, described as Hyaena variabilis by Zdansky, 1 6 3 in most of its features agrees with Crocuta eximia. That is, as has already been sug-gested by Arambourg and Piveteau 1 6 4 and Pilgrim, 1 6 5 Crocuta variabilis is only a form of the earlier established species Crocuta eximia (Roth and Wagner). Crocuta variabilis (Zdansky) differs from Crocuta eximia mainly in showing a greater variation in the anterior tubercle of the second and third upper premolars, which, as has been stated by Zdansky 1 6 6 and Pilgrim, 1 6 7 is absent in some and present in other specimens. However, these differences are not at all incompatible with a variety rank within the species Crocuta eximia (Roth and Wagner), as has been remarked by Pilgrim. 1 6 8
Considering Crocuta variabilis and Crocuta eximia as one species, the upper and lovver jaws from Kuyutarla differ from the known specimens of Crocuta eximia (Roth and Wagner), only in having a lower crown index in P1 and P2, which must, however, be regarded as individual variations.
In this connection a few words should also be said regarding the geological age of the Karacahasan beds. In a recent study, Oğuz Erol has
1 6 1 Ibid.
1 6 2 It may be pointed out here that considerable variation occurs in the anterior tubercles of the lower premolars in the M a r a g h a form of Crocuta eximia. In the specimen depicted by De M e c q u e n e m (1925, pl. I X , fig. 7) the second a n d third lower premolars are devoid of an anterior tubercle a n d in its fourth lower premolar the anterior tubercle is smaller than the posterior tubercle, as in the Pikermi form of Crocuta eximia. On the other h a n d , in the specimen depicted by Kittl (1887, pl. X V I I , fig. 1) P2 a n d P3 exhibit a small anterior tubercle a n d in P4 the anterior tubercle is larger t h a n the posterior tub ercle, as in Crocuta variabilis. T h u s in the range of variation occurring in its lower pre molars the M a r a g h a form of Crocuta eximia connects the European forms of Crocuta eximia w i t h Crocuta variabilis.
1 6 3 Zdansky, 1924, p. 93.
1 6 4 A r a m b o u r g a n d Piveteau, 1929, p p . 124-125.
1 6 5 Pilgrim, 1931, p. 117. Regarding the affinities of Crocuta variabilis, Pilgrim (1931, p. 117) states: " H y a e n a variabilis Zdansky is so closely allied to Crocuta eximia as possibly to represent a Chinese race of this species."
1 6 6 Zdansky, 1924, p. 95. 1 6 7 Pilgrim, 1931, p. 117. 1 6 8 Ibid., p. 117.
CROCUTA FROM THE KÜÇÜKYOZGAT DISTRICT 51
concluded that the whitish calcareous marls of lacustrine origin at Küçük-yozgat antedate the heterogeneous calcareous clays of running water origin at Kuyutarla and Örücebağ.1 6 9 The f a u n a1 7 0 from the whitish cal careous marls, determined to date, clearly shows that these belong to the Pontian Age,171 that is to the lower Pliocene.1 7 2 The presence of Crocuta eximia (Roth and Wagner) at Kuyutarla Locus 2, where it occurs with Hipparion gracile Kaup, shows that these stream deposits are also of Pon tian Age, representing a later horizon of the Pontian than the whitish calcareous marls of lacustrine origin.
C O N C L U S I O N
1. The remains of Crocuta found at Kuyutarla Locus 2 belong to Crocuta eximia (Roth and Wagner), which is a variable species that extended in the Pontian Age all the way from France to China.
2. The presence of Crocuta eximia (Roth and Wagner), which is a char-acteristic Pontian species, shows that the deposits of running water origin at Kuyutarla and Örücebağ belong to the Pontian, that is to the Lower Pliocene, representing a horizon of the Pontian at the Küçükyozgat district later than the whitish calcareous marls of lacustrine origin.
R E F E R E N C E S
ARAMBOURG, C. and PIVETEAU, J . 1929. Les Vertebres du Pontien de Saloniaue. Annales de Paleontologie, Vol. X V I I I , pp. 57-140.
BOULE, M. and PIVETEAU, J . 1935. Les fossiles. Elements de Paleontologie. Paris.
CHAPUT, E. 1936. Voyages d'e'tudes geologiaues et geomorphogenigues en Turguie. Memoires de I'lnstitut Français d'Archeologie de Stamboul, 11, Paris.
D E L CAMPANA, D. 1914. La Lycyaena Lunensis N. Sp. Dell'Ossario Pliocenico di Olivola (Val di Magra ). Palaeontographia Italica, Vol. XX, pp. 87-103.
D E MECQUENEM, R. 1924—1925. Contributions â l'etüde des fossiles de Maragha. Annales de Paleontologie, Vol. X I I I , 1924, pp. 133-160 and Vol. XIV, 1925, pp. 1-36.
1 6 9 Erol, 1954, pp.
93-95-1 7 0 Tschachtli, 1942, p. 326; Şenyürek, 1953a, p. 2; Şenyürek, 1953b, p. 142; Şenyürek, 1953d, p. 460; Şenyürek, 1954 b.
1 7 1 Tschachtli, 1942, p. 327; Şenyürek, 1953a, p. 2; Şenyürek, 1953b, p. 142; Şenyürek, 1953d, P- 460; Şenyürek, 1954 b.
1 7 2 As is known, Zittel (1925, p. 300), Abel (in Weber, 1928, p. XVIII), Lewis (1937, p. 194), Romer (1946, Table 4), Thenius (1949), Papp and Thenius (1949, Table VI) and Şenyürek (1951, p. 70; 1952, p. 474; 1953a, P- 2; 1953 c, p. 174; 1953d, p. 460; 1954a, p. 2) attribute the Pontian Age to the Lower Pliocene period, instead of the upper Miocene.
